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Pterygotus anglicus
Pterygotus is a genus of giant predatory eurypterid, a group of extinct aquatic arthropods. Fossils of Pterygotus have been discovered in deposits ranging in age from Late Silurian to Early Devonian, and have been referred to several different species. Fossils have been recovered from four continents; Australia, Europe, North America and South America, which indicates that Pterygotus might have had a nearly cosmopolitan (worldwide) distribution. The type species, P. anglicus, was described by Swiss naturalist Louis Agassiz in 1839. Agassiz mistakenly believed the remains were of a giant fish, with the name Pterygotus translating to "winged fish". He would only realize the mistake five years later in 1844. Pterygotus was among the largest eurypterids. Isolated fossil remains of a large chelicera (frontal appendage) suggests that the largest known species, P. grandidentatus, reached a body length of 1.75 metres (5.7 ft). Several other species, notably P. impacatus at 1.65 metres (5.4 ft) and P. anglicus at 1.6 metres (5.2 ft) were similarly gigantic. Pterygotus was surpassed in size by other giant eurypterids. Acutiramus was able to surpass 2 metres (6.6 ft), and Jaekelopterus could reach 2.6 metres (8.5 ft). Many species were considerably smaller than the largest species, such as P. kopaninensis at 50 centimetres (20 in). Like its close relative Jaekelopterus, Pterygotus was a large and active predator noted for its robust and enlarged cheliceral claws that would have allowed it to puncture and grasp prey and a visual acuity (clarity of vision) comparable to that of modern predatory arthropods. With the largest species, P. grandidentatus, reaching a body length of 1.75 metres (5.7 ft), Pterygotus was among the largest known eurypterids to have existed, though some of its close relatives (such as Acutiramus and Jaekelopterus) surpassed it in length. Though there were a few gigantic species, many species were considerably smaller in size. The smallest species, P. kopaninensis, measured just 50 centimetres (20 in) in length. Pterygotus is classified as part of the pterygotid family of eurypterids, to which it lends its name, a group of highly derived eurypterids of the Silurian to Devonian periods that differ from other groups by a number of features, perhaps most prominently in the chelicerae (the first pair of limbs) and the telson. The chelicerae of the Pterygotidae were enlargened and robust, clearly adapted to be used for active prey capture and more similar to the claws of some modern crustaceans, with well developed teeth on the claws, than to the chelicerae of other eurypterid groups. Unlike most of the rest of the body, which was covered in a scale-like ornamentation like other pterygotid eurypterids, the claws lacked any type of ornamentation. Additionally, the end points of the claws were round and curved unlike the sharp points present at the ends of the claws of the related Erettopterus. The pterygotid telsons were flattened and expanded, likely used as rudders when swimming. Their walking legs were small and slender, without spines, and they were likely not capable of walking on land. Pterygotus is distinguishable from other pterygotids by the curved distal margin of the chelae (claws). The prosoma (head) is subtrapezoidal (a trapezoid with rounded corners), with compound eyes located near the edge of the front corners. The telson (the posteriormost segment of its body) has a pronounced dorsal carina (or keel) running down its center, terminating in a short spine. Initial finds The first fossils of Pterygotus were found in deposits of Lochkovian-Pragian (Early Devonian) age by quarrymen in Scotland and western England, who referred to the large fossil remains as "Seraphims". Louis Agassiz, a Swiss-American biologist and geologist, described the fossils in 1839 and named it Pterygotus in the belief that they were the fossil remains of a large fish; "Pterygotus" translating to "winged fish". The specimens described by Agassiz from England were referred to a species he dubbed Pterygotus problematicus. Agassiz first recognized the true nature of the fossils as arthropod remains five years later in 1844 after having examined more complete fossils recovered in the Old Red Sandstone of Scotland. Although recognizing the fossils of Pterygotus as arthropod was closer to its modern phylogenetical position, Agassiz would consider Pterygotus to represent a crustacean of the Entomostraca subclass. Although Frederick M'Coy did note that Pterygotus resembled the Limulidae and the previously discovered eurypterid Eurypterus in 1849, he classified both Eurypterus and Pterygotus as crustaceans. The new Scottish fossils were named as the species P. anglicus in 1849, which remains the most extensively known species of Pterygotus, distinguished from subsequently discovered species by possessing curved terminal teeth and the primary and intermediate teeth being inclined slightly backwards. P. problematicus was also used as the designation for an incomplete chelicera discovered in the Welsh Borderland of western England by John William Salter in 1852 but is in modern times considered a nomen vanum("baseless name") as the species is impossible to define. The fossil material with which it was described is undiagnostic and insufficient to establish any meaningful characteristics and as such man… Pterygotus is classified within the family Pterygotidae in the superfamily Pterygotioidea, lending its name to both its family and its superfamily. The three most derived pterygotid eurypterids, Acutiramus, Jaekelopterus and Pterygotus, are very similar to each other. Pterygotus is particularly similar to Jaekelopterus, from which it is virtually only distinct in features of the genital appendage and potentially the telson. Similarities in the genital appendage could mean that the three genera are all synonyms of each other, as they had been classified in the past (as species of Pterygotus). Some differences between them have also been noted in the chelicerae, though chelicerae have been questioned as the basis of eurypterid generic distinction since their morphology depends on the lifestyles and has been observed to vary throughout ontogeny. Telson morphology is sometimes used as a distinguishing feature, though the telsons of the three derived pterygotid genera are all paddle-shaped (the telson of Jaekelopterus is triangular, but might fall into the morphological range of the other genera). An inclusive phylogenetic analysis with multiple species of Acutiramus, Pterygotus and Jaekelopterus is required to resolve whether or not the genera are synonyms of each other. The cladogram below is based on the nine best-known pterygotid species and two outgroup taxa (Slimonia acuminata and Hughmilleria socialis). The cladogram also contains the maximum sizes reached by the species in question, which have been suggested to possibly have been an evolutionary trait of the group per Cope’s Rule ("phyletic gigantism"). The cheliceral morphology and visual acuity of the pterygotid eurypterids separates them into distinct ecological groups. The primary method for determining visual acuity in arthropods is by determining the number of lenses in their compound eyes and the interommatidial angle (shortened as IOA and referring to the angle between the optical axes of the adjacent lenses). The IOA is especially important as it can be used to distinguish different ecological roles in arthropods, being low in modern active arthropod predators. Both Pterygotus anglicus and Jaekelopterus rhenaniae had a very high visual acuity, which researchers could determine by observing a low IOA and a large number of lenses in their compound eyes. The chelicerae of Pterygotus were enlarged, robust and possessed a curved free ramus and denticles of different lengths and sizes, all adaptations that correspond to strong puncturing and grasping abilities in extant scorpions and crustaceans. The IOA values for both Pterygotus and Jaekelopterus match those of high level and active modern predatory arthropods, indicating that they represented visual and active predators. All known pterygotids (though they are so far unknown in Ciurcopterus ) possessed cheliceral claws. The first joint of the chelicerae, where it connects to the epistoma (a plate located on the prosoma, or "head"), would have been capable of turning the entire appendage in a twisting way, which has led researchers to conclude that the function of the chelicerae would not have been only, or even primarily, for defense but rather to capture and convey food to the mouth. When captured, prey would need to be broken into smaller pieces to be able to fit into the mouth; eurypterid mouths were even less adapted to devour large pieces than mouths of modern crabs are. The eurypterid walking appendages could not cut, transport or Like Acutiramus, Pterygotus lived in fully marine environments, and its fossils are today recovered associated with common and diverse fossils indicative of such an environment. Notably, P. barrandei and P. cobbi occur in marine environments associated with fossils of Acutiramus. Pterygotus was more widespread than Acutiramus and as such many species occur without associated representatives of Acutiramus as well, such as the British P. lightbodyi. The living environment of P. australis, also marine, was in particularly deep water. The only known fossil organisms of the same age and place as P. australis are crinoids of the genus Dendrocrinus and Kooptoonocrinus and ophiouroids of the genus Protaster. The deposits where the holotype specimen of P. kopaninensis was found preserves fossils of many other animals as well. Among them are trilobites such as Leonaspis, Raphiophorus and a fragmentary harpetid trilobite. Also preserved are bivalves, such as Cardiola, and graptolites, such as Monograptus. Pterygotus most typically occurs in ecosystems with diverse eurypterid faunas, P. lanarkensis of the Kip Burn Formation of Scotland occurs together with several other eurypterid genera; Slimonia, Eusarcana, Nanahughmilleria, Parastylonurus, Erettopterus and Carcinosoma. P. impacatus from Estonia occurs together with genera Erettopterus, Erieopterus, Carcinosoma, Mixopterus, Dolichopterus and Eysyslopterus as well as cephalaspidomorph fishes. P. monroensis lived alongside Mixopterus, Hughmilleria, Eurypterus and Carcinosoma. P. nobilis lived alongside representatives of Acutiramus, Erettopterus and Eusarcana as well as with a diverse fauna of conodonts, gastropods, cephalopods, ostracods, malacostracans, trilobites and bivalves and cartilaginous fish Onchus. In the Early Devonian Dundee Formation of Scotland, fossils of P. anglicus occur together with a wide array of different eurypterids, including Erieopterus, Pagea, Tarsopterella and Stylonurus, as well as acanthodian. Category:Primeval characters Category:Sea monsters Category:Silurian creatures Category:European monsters Category:Extinct invertebrates Category:Silurian monsters of Europe Category:Prehistoric marine invertebrates Category:Eurypterids Category:Monsters